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Higher Processing of Visual Information Lecture I --- April 2, 2007 by Mu-ming Poo 1. Overview of the Mammalian Visual System 2. Retinotopic Maps and Cortical Magnification 3. Structure Lateral Geniculate Nucleus (LGN) 4. LGN Receptive Fields 5. Organization of Mammalian Visual Cortices 6. Structure of the Primary Visual Cortex Part I. Overview of Visual Pathway Projections of RGCs to the central nervous system 1. Optic nerve and optic chiasm Optic nerve consists of axons of retinal ganglion cells (RGCs). At the optic chiasm, RGC axons from nasal hemiretina cross and enter the contralateral hemisphere. RGC axons from temporal hemiretina enter the ipsilateral hemisphere. 2. Mid-brain projections RGC axons project to pretectum, which controls pupillary reflex. RGC axons project to superior colliculus, which controls saccadic eye movement. 3. Central projections LGN axons project to the primary visual cortex (striate cortex, V1) on the same side Retinotopic map All structures in the geniculo-cortical system have retinotopic maps. Every point on the map corresponds to a point on the retina. Cortical magnification In cortex, the area devoted to the foveal region of the retina is enlarged relative to the rest of the eye. Cortex Visual field LGN Visual field Fovea: 1 cone => 1 RGC 1 RGC => 2-3 LGN cells 1 LGN cell => 1000 V1 cells 10o periphery: 2 cone => 1 RGC 2 RGCs => 1 LGN cell 1 LGN cell => 100 V1 cells Cortex Structure of the primate LGN Each LGN has 6 layers labeled 1 thru 6 from ventromedial to dorsolateral. 6 5 Magnocellular LGN layers - Layers 1 & 2 - inputs from M-type ganglion cells; 4 Parvocellular LGN layers - Layers 3,4,5,6 - inputs from P-type ganglion cells; 3 2 1 Koniocellular LGN layers – Interlayers just ventral to each numbered layer - inputs from blue/yellow ganglion cells 50m Layers 1,3,6 receive contralateral input and Layers 2,3,5 receive ipsilateral input. Each LGN serves the contralateral visual field. The retinotopic maps from two eyes are in register across the layers. (Axons from RGCs responding to the same visual field innervate LGN cells that are aligned vertically across the layers) Properties of P, M, and K cells Cortical Feedback to LGN: The corticogeniculate pathway Only <30% inputs to LGN are from the retina. >70% are feedback inputs, mostly from V1. The corticogeniculate axons terminate in all LGN layers in two main patterns (Ichida & Casagrande, 2002) Common functional patterns of corticogeniculate connectivity (Sillito et al, Science 1999): (1) in individual M or P LGN layers. Feedback to LGN cells along the preferred orientation, enhancing the buildup of orientation selectivity within the geniculo-cortical circuit. Feedback to LGN cells along the axis of stimulus movement, enhancing responsiveness in anticipation of stimulus movement. (2) Terminate in pairs of M, P, or K layers. Receptive Fields of RGCs Leaving the retina ON-OFF Center-surround receptive field Color opponency M cells P cells B/Y (K) cells 3. LGN receptive fields Each LGN neuron receives input from only a few ganglion cells. The receptive fields of LGN neurons are concentric with a far stronger surround antagonism than that in RGCs, suggesting the LGN is not a simple relay station. (Hubel & Wiesel, 1962) Brodmann’s cytoarchitectonic maps of cerebral cortex Side view of left hemisphere Inside view of right hemisphere from the midline The organization of mammalian visual cortices monkey monkey human Anatomy of the primary visual cortex (V1, striate cortex) V1 has six layers Layer 1 is nearly aneuronal, composed predominantly of dendritic and axonal connections. Layers 2/3 contain excitatory neurons which project to extrastriate cortical regions Layer 4 is divided into 4A, 4B, 4Ca, and 4Cb. Layers 4Ca and 4Cb are the major recipients of LGN projections. Layers 5/6 contain excitatory projection neurons which provide feedback to LGN. Blobs --Cytochrome oxidase (CO) labeling of V1 shows CO rich regions in layer II & III termed "blobs" Layer 3 Cortical cell types Excitatory cells: Spiny pyramidal & stellate cells Inhibitory cells: >40 types Inputs from LGN Most LGN cells terminate in layer 4, M in 4Ca, P mostly in 4Cb. Collaterals (branches) of M & P terminate in 6. K cells and some P cells terminate in blobs in layers 2 & 3. Output from V1 Layers II, III & IVB - feedward to extrastriate cortical areas Layer V - feedback to pons and superior colliculus Layer VI - feedback to LGN